Mycosystema
菌 物 学 报
15 January 2009, 28(1): 001-013
[email protected]
ISSN1672-6472 CN11-5180Q
©2009 Institute of Microbiology, CAS, all rights reserved.
A new systematic arrangement of the genus
Oudemansiella s. str. (Physalacriaceae, Agaricales)
YANG Zhu-Liang1*
ZHANG Li-Fang1
Gregory M. MUELLER2
Gerhard W. KOST3
Karl-Heinz REXER3
1
Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China
2
Department of Botany, Field Museum of Natural History, Chicago, IL 60605-2496, U.S.A.
3
Systematic Botany & Mycology, FB17, Philipps-University Marburg, 35032 Marburg, Germany
Abstract: The genus Oudemansiella s. str. has been revised and circumscribed as an assemblage of taxa excluding the genus Xerula
s. str. A new systematic arrangement with four sections, i.e., Oudemansiella, Mucidula, Dactylosporina and Radicatae, is proposed.
Section Oudemansiella is comprised of tropical to south temperate species, e.g., O. platensis, O. australis, O. canarii and O.
crassifolia, with an ixotrichoderm pileipellis composed of filamentous hyphae that, in many cases, are intermixed with inflated cells
often occurring in chains, while section Mucidula encompasses north temperate and subtropical taxa such as O. mucida, O.
venosolamellata and O. submucida that have an ixohymeniderm-trichoderm pileipellis composed of more or less clavate terminal
cells. Sections Oudemansiella and Mucidula share similar habitats and forms of basidiomata, i.e., with or without a (rudimentary)
annulus on the stipe and growing on exposed rotten wood. Section Dactylosporina accommodates species from South and Central
America with basidiospores that have finger-like ornamentation. Section Radicatae, represented by O. radicata and its allies, is the
largest section and includes the remaining species of the genus in its restricted sense. Oudemansiella americana, O. caussei and O.
hongoi, previously treated in O. sect. Albotomentosae or Xerula sect. Hyalosetae, are excluded from Oudemansiella in this treatment
because preliminary data indicate that they represent a segregate genus. Section Mucidula, 32 combinations and 1 name are newly
proposed.
Key words: Basidiomycetes, new combinations, new name, nomenclature, systematics, taxonomy
狭义小奥德蘑属（膨瑚菌科,蘑菇目）的一个新系统
杨祝良
1
张丽芳
Karl-Heinz REXER
1
Gregory M. MUELLER
2
3
Gerhard W. KOST
3
Supported by the National Natural Science Foundation of China (No. 30525002, 30470010 and 30700005), the Knowledge Innovation Program of the
Chinese Academy of Sciences (No. KSCX2-YW-G-025), the National Basic Research Program of China (No. 2009CB522300) and the Joint Funds of
the National Natural Science Foundation of China and Yunnan Provincial Government (No. U0836604). Part of the field work was supported by the
National Science Foundation of the USA (DEB-0321846 to D.E. Boufford and Richard H. Ree).
*
Corresponding author. E-mail: [email protected]; [email protected]
Received: 19-12-2008, accepted: 22-12-2008
Mycosystema
2
1
中国科学院昆明植物研究所生物多样性与生物地理学重点实验室 昆明 650204
2
美国芝加哥费尔德自然历史博物馆植物部 芝加哥 IL 60605-2496
3
德国马尔堡大学生物学系系统植物学与真菌学实验室 马尔堡 35032
摘
要：本文对狭义小奥德蘑属 Oudemansiella s. str.的概念做了修订，在修订后的属中，狭义干蘑属 Xerula s. str.的物种不
纳入其中。在狭义小奥德蘑属下，提出了一个包含 4 个组 O. sect. Oudemansiella、Mucidula、Dactylosporina 和 Radicatae
的新系统。小奥德蘑组 sect. Oudemansiella 包括热带至南温带的一些物种，如新热带小奥德蘑 O. platensis、澳洲小奥德蘑
O. australis、旧热带小奥德蘑 O. canarii 和宽褶小奥德蘑 O. crassifolia，这些物种的菌盖表皮为粘栅栏型，由菌丝组成，但其
中常夹杂有链状排列的膨大细胞。粘蘑组 sect. Mucidula 包含北半球温带至亚热带的一些物种，如粘小奥德蘑 O. mucida、网
褶小奥德蘑 O. venosolamellata 和近粘小奥德蘑 O. submucida，其菌盖表皮为粘子实层-栅栏型，由近棒状的顶端膨大细胞组
成。小奥德蘑组和粘蘑组的物种，在外形和小生境上有相似之处，其担子果皆生于地表外的腐木上，菌柄上有或无菌环。刺
孢组 sect. Dactylosporina 包含中南美洲那些孢子表面有指状凸起的物种。长根组 sect. Radicatae 由长根小奥德蘑 O. radicata
及其近缘种为代表，是该属中最大的组，包括该属其他三组之外的所有种。北美的 O. americana、欧洲的 O. caussei 和东亚
的 O. hongoi 曾被置于小奥德蘑属中的白毛组 O. sect. Albotomentosae 或干蘑属的亮毛组 X. sect. Hyalosetae，在本系统中它们
没有纳入小奥德蘑属，因为它们可能代表一个单独的属。本文还提出了 1 新等级、32 个新组合和 1 个新名称。
关键词：担子菌，新组合，新名称，命名法，系统学，分类学
INTRODUCTION
Petersen 2000; Mueller et al. 2001; Petersen & Hughes
2005; Horak 2005; Petersen & Nagasawa 2006; Petersen
The genus Oudemansiella Speg. was proposed by
& Baroni 2007; Petersen 2008a, 2008b, 2008c). Among
Spegazzini (1881) based on the Neotropical Agaricus
the contributions to the study of Oudemansiella-Xerula,
platensis Speg. Mucidula Pat. (1887) initially included
Dörfelt’s work has had great influence. He greatly
only A. mucidus Schrad.: Fr. from Europe. Boursier
emended the genus Xerula by transferring sections
(1924) emended Mucidula to include A. longipes Bull.
Albotomentosae Clémençon, Protoxerula Clémençon
and A. radicatus Relhan. The genus Xerula Maire was
and Radicatae Clémençon from Oudemansiella to
proposed by Maire (1933) based on A. longipes.
Xerula, and restricting Oudemansiella only to species
Moser (1955) merged Xerula and Mucidula into
with a (rudimentary) annulus, such as O. mucida and O.
Oudemansiella. Singer (1962a, 1962b, 1964, 1986)
canarii (Dörfelt 1979, 1980a, 1980b, 1981, 1983, 1984).
adapted Moser’s treatment, yet regarded Xerula as a
His concept was followed by other authors, and many
subgenus within the genus Oudemansiella. Clémençon
species have subsequently been transferred from
(1979) provided a classification of Oudemansiella and
Oudemansiella to, or newly described in, Xerula. Several
treated Xerula as one of five subgenera within
section
Oudemansiella based on a computer-aided study of 16
Dörfelt, Hyalosetae Dörfelt, Radicatae and Xerula have
morphological characters, while Pegler & Young (1987)
been erected in Xerula which has been divided into three
divided Oudemansiella into five sections under two
subgenera,
subgenera, Oudemansiella and Xerula. The treatment of
Ixoflammula Contu (Dörfelt 1980b; Contu 2000).
names,
viz.
e.g.,
Albotomentosae
Xerula,
Radicatae
(Clémençon)
Dörfelt
and
Pegler & Young (1987) was adopted by Rexer & Kost
Recent molecular phylogenetic analyses place
(1989a, b), Yang & Zang (1993), Yang (2000) and
Oudemansiella and Xerula in the Physalacriaceae clade
Mizuta (2006).
(Moncalvo et al. 2002; Wilson & Desjardin 2005;
Some mycologists retained Oudemansiella and
Matheny et al. 2007), and document that taxa of
Xerula as two separate genera (Dörfelt 1979; Boekhout
Oudemansiella and Xerula are resolved into two separate
& Bas 1986; Redhead et al. 1987; Petersen & Halling
monophyletic clades (Zhang et al. 2003; Binder et al.
1993; Petersen & Methven 1994; Corner 1994, 1996;
2006; Zhang 2006). Petersen (2008c) stated that Xerula
Boekhout 1999; Halling & Mueller 1999; Contu 2000;
in his definition is not monophyletic. Wang et al. (2008)
Vol.28 No.1
3
treated Xerula as a distinct genus, but in a restricted
45 (HKAS 43457).
sense that includes only hispid species such as X. pudens
O. aff. submucida Corner – China. Mt. Changbai,
(Pers.) Singer and its close allies that have a collybioid
Antu County, Jilin Prov., alt. 840m, 15 August 2004, L.F.
basidioma with dry surfaces of the pileus and the stipe
Zhang 497 (HKAS 7622).
covered with erect, lanceolate, yellow to brown,
thick-walled
the
based on fresh material, colored photos, and/or
while
field-notes. For micromorphological studies, sections
Oudemansiella sensu stricto accommodates non-hispid
were cut with a razor blade from basidioma and mounted
species that have sterile lamellar edges with crowded
on slides in 5% KOH, and then observed, measured and
cheilocystidia and clavate to narrowly basidioles in the
illustrated
hymenium. Based on morphology and the molecular
Axioskop 40, Germany). Herbaria are abbreviated
phylogenetic analyses of Zhang et al. (2003) and Zhang
according to Holmgren et al. (1990) with one exception:
(2006), a new systematic arrangement for Oudemansiella
HKAS = Herbarium of Cryptogams, Kunming Institute
s. str. is proposed.
of Botany of the Chinese Academy of Sciences, which is
hymenium
setae,
and
a
subacerose
fertile
basidioles
lamellar
edge,
in
Macromorphological characters were described
1 MATERIAL AND METHODS
Over 150 collections of Oudemansiella and Xerula
under
a
compound
microscope (Zeiss
not listed in the index or related publications.
2 RESULTS AND DISCUSSIONS
from Asia, Europe, Australia, and North and Central
Morphological studies document that the structure
America were examined, but only specimens from which
of the pileipellis of O. canarii-O. platensis is much more
illustrations in the present paper were made are listed
diverse
below.
venosolamellata. In specimens of O. canarii-O. platensis,
and
complicated
than
in
O.
mucida-O.
Oudemansiella canarii (Jungh.) Höhn. – China.
the pileipellis is an ixotrichoderm composed of both
Wild Elephant Valley, Jinghong County, Yunnan Prov.,
inflated cells and branching filamentous hyphae,
alt. 650m, 1 September 2004, X.H. Wang 1820 (HKAS
sometimes dominantly of filamentous hyphae with few
5021).
inflated elements (Figs. 1, 3, 4; see also figures 4, 7 in
O. aff. crassifolia Corner – China. Longdao, Ruili
Corner 1994 and figure 8 in Baroni & Ortiz 2002, and
County, Yunnan Prov., alt. 1200m, 2 August 2003, P.C.
Petersen et al. 2008), while the pileipellis in the O.
Liu 87 (HKAS 43500).
mucida-O. venosolamellata complex, is usually a regular
O. mucida (Schrad.: Fr.) Höhn. – Germany.
ixohymeniderm
or
an
ixohymeniderm-trichoderm
Oberjoch, Allgäu, Bayern, 1 October 1997, Z.L. Yang
composed of broadly clavate, clavate to narrowly clavate
2111 (HKAS 31558).
cells in a layer of 1-2(3) cells (Figs. 6-8). The pileipellis
O. platensis (Speg.) Speg. – Colombia. Estación
of O. mucida is somewhat special because it made up of
Cientifica de la Universidad National, Mpio de Leticia,
clavate to subcylindric to irregularly or coralloidly
Dpto de Amazonas, 4 May 1992, A.E. Franco-Molano
branched terminal cells (Fig. 5; also see Petersen et al.
810 (HKAS 45442, duplicate of NY 34703).
2008). In addition, there are often flecks or warts on the
O. venosolamellata (Imazeki & Toki) Imazeki &
pileal surface in specimens of O. canarii-O. platensis
Hongo – Japan. Mt. Hayachine, Kawai, Iwate Prefecture,
(Fig. 2) but not in the O. mucida-O. venosolamellata
8 August 1988, H. Neda s.n. (TFM-M-A843).
complex. The pileipellis of O. radicata complex is
O. steffenii (Rick) Singer – Brazil. São Leopoldo,
usually a regular ixohymeniderm composed of broadly
Rio Grande do sul, 1932, J. Rick s.n. [FH, holotype of
clavate, clavate to narrowly clavate cells. A few species
Oudemansiella echinosperma Singer, which is usually
of this complex have no pileocystidia (Fig. 13a of Rexer
treated as a synonym of O. steffenii by Singer (1964) and
& Kost 1989a), somewhat similar to those of O.
other authors].
mucida-O. venosolamellata complex, but most possess
O. submucida Corner – China. Mengxiu, Ruili
pileocystidia (Figs. 5-6 of Yang 2000; Fig. 11 of Yang &
County, Yunnan Prov., alt. 1190m, 28 July 2003, P.C. Liu
Zhang 2003). The pileipellis of O. steffenii (Fig. 9) is
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Mycosystema
4
very similar to those of O. radicata complex but without
pileocystidia.
Fig. 1 Radial section of pileipellis from median between pileal centre and margin of O. platensis (HKAS 45442); Fig. 2 Vertical section of a wart
on the pileal surface of O. platensis (HKAS 45442); Fig. 3 Radial section of pileipellis from median between pileal centre and margin of O.
canarii (HKAS 5021); Fig. 4 Radial section of pileipellis from median between pileal centre and margin of O. aff. crassifolia (HKAS 43500); Fig.
5 Radial section of pileipellis from median between pileal centre and margin of O. mucida (HKAS 31558). Scale bar = 20μm.
(1881).
3 TAXONOMY
Syn.: Oudemansia Speg., Anal. Soc. Cient. Argent. 10:
The structure of the pileipellis in Oudemansiella is
very diverse and useful for the establishment of
infrageneric
delimitations.
Four
groups
can
be
recognized based on the pileipellis structure. These
groups correspond to the following four sections.
Oudemansiella Speg., Anal. Soc. Cient. Argent. 12: 24
280 (1880).
Mucidula Pat., Hymén. Eur.: 95 (1887).
Phaeolimacium Henn., in Warburg, Monsunia 1: 14
(1899).
Basidioma armillarioid, tricholomatoid, collybioid
to marasmioid. Surface of pileus usually viscid to gelatinous.
Vol.28 No.1
5
Fig. 6 Radial section of pileipellis from median between pileal centre and margin of O. venosolamellata (TFM-M-A843); Fig. 7 Radial section of
pileipellis from median between pileal centre and margin of O. aff. submucida (HKAS 7622); Fig. 8 Radial section of pileipellis from median
between pileal centre and margin of O. submucida (HKAS 43457); Figs. 9-10 Pileipellis of O. steffenii from median between pileal centre and margin,
and basidiospores O. steffenii (FH, holotype of O. echinosperma). Scale bar = 20μm.
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Mycosystema
6
Lamellae adnexed to adnate, often with short decurrent
Section 2 Mucidula (Pat.) Zhu L. Yang, Li F. Zhang,
teeth, thick, subdistant to distant. Stipe central to
G.M. Muell., G. Kost & Rexer, stat. nov.
eccentric, annulate or exannulate, with or without a
Basionym: Mucidula Pat., Hymén. Eur.: 95 (1887).
pseudorhiza. Spore print white to cream-coloured.
MycoBank No. MB 512737
Pileipellis more or less an ixohymeniderm, ixotricho-
Basidioma tricholomatoid to armillarioid. Stipe
hymeniderm or ixotrichoderm composed of inflated cells
often without a pseudorhiza, but with a permanent,
and/or filamentous hyphae. Warts or flecks on pileal
ephemeral, or fugacious annulus. Pileipellis a continuous
surface, when present, composed of more or less
and regular ixotrichohymeniderm or ixohymeniderm
vertically arranged moniliform to subglobose inflated
composed of clavate, narrowly clavate to broadly clavate,
cells intermixed with filamentous hyphae. Pileocystidia
often branched sometimes coralloid terminal cells
(hairs) present or absent, usually unicellular. Basidia
(Figs. 5-8). Flecks or patches on pileal surface absent.
mostly clavate to broadly clavate, sometimes sub-
Basidiospores globose to subglobose, sometimes broadly
urniform, often with a distinctly narrowed base;
ellipsoid or subamygdaliform to limoniform, without
basidioles in the hymenium usually clavate and with a
digitate ornamentation. In north temperate and subtro-
broadly rounded apex, rarely subacerose. Basidiospores
pical regions, sometimes also in tropical mountains.
smooth to very delicately puckered or punctulate,
Type species: O. mucida.
sometimes with digitate ornamentation, colorless and
Additional species: O. submucida, O. yunnanensis
hyaline, non-amyloid, non-dextrinoid, acyanophilic.
Zhu L. Yang & M. Zang, O. venosolamellata.
Cheilocystidia crowded and forming a sterile lamellar
Section 3 Radicatae Clémençon, Sydowia 32: 78 (1979).
edge. Clamp connections usually present and common,
= Oudemansiella sect. Pseudoradicatae Clémençon,
but for 2-spored taxa usually absent.
Sydowia 32: 77 (1979).
Type species: O. platensis.
= Oudemansiella sect. Hygrophoroides Clémençon,
Sydowia 32: 78 (1979).
Section 1 Oudemansiella
= Phaeolimacium Henn., in Warburg, Monsunia 1: 14
(1899).
≡ Xerula sect. Radicatae (Clémençon) Dörfelt,
Feddes Repert. 91: 433 (1980).
Basidioma
to
Basidioma collybioid. stipe without annulus, often
marasmiod. Stipe with a rudimentary or fugacious
but not always with a pseudorhiza. Pileipellis a
annulus,
continuous and regular ixohymeniderm composed of
often
armillarioid,
without
tricholomatoid
pseudorhiza.
Pileipellis
an ixotrichoderm, but often stretched and extensively
narrowly
distorted and, thus, irregularly arranged due to the
sphaeropedunculate terminal cells, with or without
expansion of the pileus, composed of moniliform,
pileocystidia. Basidiospores globose, ovoid, broadly
fusiform to ellipsoid to subglobose inflated and
ellipsoid to ellipsoid or amygdaliform, without digitate
filamentous hyphae (Figs. 1, 3 and 4). When basidioma
ornamentation. Widely distributed.
is fully or over mature, pileipellis often distorted and
cutis-like. Flecks or warts on pileal surface, when present,
clavate,
clavate
to
broadly
clavate
to
Type species: O. radicata (Relhan: Fr.) Singer (≡
Agaricus radicatus Relhan: Fr.)
composed of more or less vertically arranged moniliform
Additional taxa: O. bispora (Natarajan & Purush.)
to subglobose inflated cells intermixed with filamentous
Zhu L. Yang & Li F. Zhang, O. japonica (Dörfelt) Pegler
hyphae or mainly with filamentous hyphae (Fig. 2).
& Young, O. orientalis Zhu L. Yang, O. raphanipes
Basidiospores globose to subglobose, without digitate
(Berk.) Pegler & T.W.K. Young, and the following taxa.
ornamentation. Tropical, subtropical to the south
Oudemansiella africana (Dörfelt) Zhu L. Yang, G.M.
temperate.
Muell., G. Kost & Rexer, comb. nov.
Type species: O. platensis.
Basionym: Xerula radicata var. africana Dörfelt, Feddes
Additional species: O. canarii, O. australis G. Stev.
Repert. 95: 195, Abb. 2, Taf. 23/10-12 (1984).
& G.M. Taylor, O. crassifolia, O. fibrillosa T.J. Baroni &
B. Ortiz, O. lianicola Corner.
≡ Xerula africana (Dörfelt) R.H. Petersen, Fungal
Diversity 30: 122 (2008).
Vol.28 No.1
7
≡ Oudemansiella radicata var. africana (Dörfelt)
≡ Oudemansiella japonica var. colensoi (Dörfelt)
Pegler & T.W.K. Young, Trans. Brit. Mycol. Soc. 87: 598
Pegler & T.W.K. Young, Trans. Brit. Mycol. Soc. 87: 596,
(1987).
figs. 11, 52 (1987).
MycoBank No. MB 512738
≡ Xerula colensoi (Dörfelt) R.H. Petersen, Nova
Oudemansiella alveolata (Kalchbr.) Zhu L. Yang, G.M.
Muell., G. Kost & Rexer, comb. nov.
Hedwigia 87: 9, figs. 11-16 (2008).
MycoBank No. MB 512749
Basionym: Agaricus alveolatus Kalchbr., Grevillea 9:
Oudemansiella crassibasidiata (R.H. Petersen) Zhu L.
110 (1881).
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
≡ Collybia alveolata (Kalchbr.) Sacc., Syll. Fung. 5:
202 (1887).
Basionym: Xerula crassibasidiata R.H. Petersen, Fungal
Diversity 30: 128, figs. 12-17 (2008).
≡ Xerula alveolata (Kalchbr.) R.H. Petersen, Fungal
Diversity 30: 125, figs. 7-11 (2008).
MycoBank No. MB 512750
Oudemansiella eradicata (Kalchbr.) Zhu L. Yang, G.M.
MycoBank No. MB 512739
Muell., G. Kost & Rexer, comb. nov.
Oudemansiella amygdaliformis var. bispora (R.H.
Basionym: Agaricus (Collybia) eradicatus Kalchbr.,
Petersen & Nagas.) Zhu L. Yang, G.M. Muell., G. Kost &
Grevillea 8: 151 (1880).
Rexer, comb. nov.
Basionym: Xerula amygdaliformis var. bispora R.H.
Petersen & Nagas., Rep. Tottori Mycol. Inst. 43: 12, figs.
6-8 (2006).
Oudemanisella atrocaerulea (R.H. Petersen & Bougher)
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Basionym: Xerula atrocaerulea R.H. Petersen &
Bougher, Nova Hedwigia 87: 5, figs. 1, 2, 5-10 (2008).
MycoBank No. MB 512743
Oudemansiella aureocystidiata (R.H. Petersen & Nagas.)
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Basionym: Xerula aureocystidiata R.H. Petersen &
Nagas., Rep. Tottori Mycol. Inst. 43: 14, figs. 9-14
(2006).
Oudemansiella caulovillosa (R.H. Petersen) Zhu L.
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
caulovillosa
≡ Xerula eradicata (Kalchbr.) R.H. Petersen, Nova
= Agaricus (Collybia) olivaceo-albus Cooke & Massee,
Grevillea 15: 93 (1887); Collybia olivaceo-albus (Cooke
& Massee) Sacc. Syll. Fung. 9: 28 (1891). (fide Petersen
2008).
MycoBank No. MB 512751
Oudemansiella flavo-olivacea (R.H. Petersen & Halling)
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Basionym: Xerula flavo-olivacea R.H. Petersen &
Halling, Nova Hedwigia 87: 19, figs. 23-30 (2008).
MycoBank No. MB 512752
Oudemansiella flavo-olivacea var. kimberleyana (R.H.
MycoBank No. MB 512746
Xerula
202 (1887).
Hedwigia 87: 15, figs. 17-22 (2008)
MycoBank No. MB 512741
Basionym:
≡ Collybia eradicata (Kalchbr.) Sacc., Syll. Fung. 5:
R.H.
Petersen,
Mycoscience 49: 26, figs. 53-64 (2008).
MycoBank No. MB 512783
Oudemansiella chiangmaiae (R.H. Petersen & Nagas.)
Petersen & M.D. Barrett) Zhu L. Yang, G.M. Muell., G.
Kost & Rexer, comb. nov.
Basionym: Xerula flavo-olivacea var. kimberleyana H.
Petersen & M.D. Barrett, Nova Hedwigia 87: 23, figs.
31-38 (2008).
MycoBank No. MB 512753
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella furfuracea (Peck) Zhu L. Yang, G.M.
Basionym: Xerula chiangmaiae R.H. Petersen & Nagas.,
Muell., G. Kost & Rexer, comb. nov.
Rep. Tottori Mycol. Inst. 43: 17, figs. 15-20 (2006).
Basionym: Collybia radicata var. furfuracea Peck,
MycoBank No. MB 512748
Oudemansiella colensoi (Dörfelt) Zhu L. Yang, G.M.
Muell., G. Kost & Rexer, comb. nov.
Report. N. Y. State Mus. 45 (Bot. Ed.): 31(Mus. Ed. P.
91) (1892).
≡ Oudemansiella radicata var. furfuracea (Peck)
Basionym: Xerula japonica var. colensoi Dörfelt, Feddes
Pegler & T.W.K. Young, Trans. Brit. Mycol. Soc. 87:
Repert. 95: 193 (1984).
598 (1987).
http://journals.im.ac.cn/jwxtcn
Mycosystema
8
≡ Xerula furfuracea (Peck) Redhead, Ginns &
Shoemaker, Mycotaxon 30: 362 (1987).
MycoBank No. MB 512754
Oudemansiella gigaspora (Cooke & Massee) Zhu L.
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Basionym: Clitocybe megalospora Clem., Bot. Survey
Nebraska 4: 18 (1896).
≡ Xerula megalospora (Clem.) Redhead, Ginns &
Shoemaker, Mycotaxon 30: 374 (1987).
MycoBank No. MB 512762
Basionym: Hygrophorus (Camarophyllus) gigasporus
Oudemansiella mundroola (Grgur.) Zhu L. Yang, G.M.
Cook & Massee, Grevillea 16: 31 (1887).
Muell., G. Kost & Rexer, comb. nov.
≡ Xerula gigasporus (Cook & Massee) H. Petersen,
Nova Hedwigia 87: 27, figs. 41-43 (2008).
= Xerula radicata var. australis Dörfelt, Feddes Repert.
95: 195, Taf. 23/9, 13-14 (1984), non Oudemansiella
australis G. Stev. & G.M. Taylor, Kew Bull. 19: 33
Basionym: Xerula radicata var. mundroola Grgur.,
Larger Fungi of South Australia (Adelaide): 253 (1997).
≡ Xerula mundroola (Grgur.) R.H. Petersen, Nova
Hedwigia 87: 43, figs. 39,40, 44-46 (2008).
MycoBank No. MB 512763
(1964); Oudemansiella radicata var. australis (Dörfelt)
Oudemansiella orientiradicata Zhu L. Yang, G.M.
Pegler and T.W.K. Young, Trans. Brit. Mycol. Soc. 87:
Muell., G. Kost & Rexer, nom. nov.
598 (1987); Xerula australis (Dörfelt) R.H. Petersen,
Basionym: Xerula orientalis R.H. Petersen & Nagas.,
Can. J. Bot. 72: 1152 (1994).
Rep. Tottori Mycol. Inst. 43: 36, figs. 46-52 (2006); non
MycoBank No. MB 512755
Oudemansiella globospora (R.H. Petersen & Nagas.)
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella orientalis Zhu L. Yang, Mycotaxon 74:
357 (2000).
MycoBank No. MB 512765
Basionym: Xerula globospora R.H. Petersen & Nagas.,
Oudemansiella orientiradicata var. margaritella (R.H.
Rep. Tottori Mycol. Inst. 43: 25, figs. 28-34 (2006).
Petersen & Nagas.) Zhu L. Yang, G.M. Muell., G. Kost
MycoBank No. MB 512757
& Rexer, comb. nov.
Oudemansiella incognita (Methven & R.H. Petersen)
Basionym: Xerula orientalis var. margaritella R.H.
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Petersen & Nagas., Rep. Tottori Mycol. Inst. 43: 39, fig.
Basionym: Xerula incognita Methven & R.H. Petersen,
53 (2006).
Can. J. Bot. 72: 1154, figs. 5-9 (1994).
MycoBank No. MB 512758
MycoBank No. MB 512767
Oudemansiella radicata var. bispora (Redhead, Ginns
Oudemansiella kenyae (R.H. Petersen) Zhu L. Yang,
& Shoemaker) Zhu L. Yang, G.M. Muell., G. Kost &
G.M. Muell., G. Kost & Rexer, comb. nov.
Rexer, comb. nov.
Basionym: Xerula kenyae R.H. Petersen, Fungal
Basionym: Xerula radicata var. bispora Redhead, Ginns
Diversity 30: 132, figs. 18-23 (2008).
& Shoemaker, Mycotaxon 30: 398, figs. 14, 20 (1987).
MycoBank No. MB 512759
MycoBank No. MB 512768
Oudemansiella limonispora (R.H. Petersen) Zhu L.
Oudemansiella rubrobrunnescens (Redhead, Ginns &
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Shoemaker) Zhu L. Yang, G.M. Muell., G. Kost & Rexer,
Basionym: Xerula limonispora R.H. Petersen, Bull. Soc.
comb. nov.
Mycol. France 120: 38 (2005).
Basionym: Xerula rubrobrunnescens Redhead, Ginns &
MycoBank No. MB 512760
Oudemansiella mammicystis (R.H. Petersen) Zhu L.
Shoemaker, Mycotaxon 30: 384, figs.4-5, 40-51 (1987).
MycoBank No. MB 512769
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella rugosoceps (G.F. Atk.) Zhu L. Yang,
Basionym: Xerula mammicystis R.H. Petersen, Fungal
G.M. Muell., G. Kost & Rexer, comb. nov.
Diversity 30: 135, figs. 24-29 (2008).
Basionym: Collybia rugosiceps G. F. Atk. Journ. Mycol.
MycoBank No. MB 512761
Oudemansiella megalospora (Clem.) Zhu L. Yang, G.M.
Muell., G. Kost & Rexer, comb. nov.
8: 112 (1902)
≡ Gymnopus rugosoceps (G.F. Atk.) Murrill, N.
Amer. Fl. (New York) 9: 360 (1916).
Vol.28 No.1
9
≡ Xerula rugosoceps (G.F. Atk.) Redhead, Ginns &
Shoemaker, Mycotaxon 30: 386 (1987).
MycoBank No. MB 512770
MycoBank No. MB 512775
Oudemansiella trichofera (R.H. Petersen) Zhu L. Yang,
G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella semiglabripes (R.H. Petersen) Zhu L.
Basionym: Xerula trichofera R.H. Petersen, Nova
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Hedwigia 87: 54, figs. 54, 61-67 (2008).
Basionym: Xerula semiglabripes R.H. Petersen, Fungal
Diversity 30: 138, figs. 30-35 (2008).
MycoBank No. MB 512771
MycoBank No. MB 512776
Oudemansiella variabilis (R.H. Petersen) Zhu L. Yang,
G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella superbiens (Berk.) Zhu L. Yang, G.M.
Basionym: Xerula variabilis R.H. Petersen, Nova
Muell., G. Kost & Rexer, comb. nov.
Hedwigia 87: 59, figs. 68-73 (2008).
MycoBank No. MB 512777
Basionym: Agaricus radicatus var. superbiens Berk.,
Hooker’s Lond. J. Bot. 4: 43 (1845).
≡ Collybia radicata var. superbiens (Berk.) Sacc.,
Syll. Fung. 5: 201 (1887).
≡ Oudemansiella radicata var. superbiens (Berk.)
Pegler & T.W.K. Young, Trans. Br. mycol. Soc. 87(4):
598 (1987).
≡ Xerula radicata var. superbiens (Berk.) Dörfelt,
Feddes Repert. 94: 559 (1983).
≡ Xerula superbiens (Berk.) R. H. Petersen, Nova
Hedwigia 87: 50, figs. 53, 55-60 (2008).
MycoBank No. MB 512773
Oudemansiella tetrasperma (R.H. Petersen) Zhu L.
Yang, G.M. Muell., G. Kost & Rexer, comb. nov.
Oudemansiella vinocontusa (R.H. Petersen & Nagas.)
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb.
nov.
Basionym: Xerula vinocontusa R.H. Petersen & Nagas.,
Rep. Tottori Mycol. Inst. 43: 44, figs. 60-64 (2006).
MycoBank No. MB 512779
Section 4 Dactylosporina (Clémençon) Pegler & T.W.K.
Young, Trans. Br. Mycol. Soc. 87: 598 (1987).
Basionym:
Oudemansiella
subgen.
Dactylosporina
Clémençon, Sydowia 32: 77 (1979).
≡ Dactylosporina (Clémençon) Dörfelt, Feddes
Repert. 96: 236 (1985).
Basidioma collybioid. Stipe without an annulus, but
Basionym: Xerula tetrasperma R.H. Petersen, Fungal
usually with a pseudorhiza. Pileipellis a continuous
Diversity 30: 141, figs. 36-40 (2008).
and regular hymenioderm composed of clavate, narrowly
MycoBank No. MB 512774
clavate to broadly clavate or sphaeropedunculate
Oudemansiella tetrasperma f. marginata (R.H. Petersen)
terminal cells, without pileocystidia (Fig. 9). Basidio-
Zhu L. Yang, G.M. Muell., G. Kost & Rexer, comb.
spores with obtusely echinate ornamentation (Fig. 10). In
nov.
South and Central America.
Basionym: Xerula tetrasperma f. marginata R.H.
Type species: O. steffenii.
Petersen, Fungal Diversity 30: 143 (2008).
Additional species: O. macracantha Singer.
Key to the sections
1. Basidiomata armillarioid or tricholomatoid, growing directly on wood above the ground, often with ephemeral or persistent
annulus; pileipellis an ixotrichoderm, ixotrichohymeniderm or ixohymeniderm with both inflated and branched filamentous hyphae
or clavate terminal elements but can become cutis-like when fully mature..............................................................................................2
1. Basidiomata collybioid, growing on buried wood, without annulus, usually with distinct pseudorhiza; pileipellis a continuous and
regular ixohymeniderm..............................................................................................................................................................................3
2. Pileipellis an ixotrichoderm, or when fully mature completely distorted and cutis-like, consisting of both filamentous hyphae and
inflated cells, sometimes primarily composed of filamentous hyphae; pileal surface often with flecks or patches; basidiospores
globose to subglobose; tropical, subtropical to south temperate........................................................................... O. sect. Oudemansiella
2. Pileipellis a regular ixohymeniderm or an ixotrichohymeniderm composed of more or less clavate terminal cells; pileal surface
without flecks or patches; basidiospores globose to subglobose, sometimes broadly ellipsoid or subamygdaliform to limoniform;
north temperate and subtropical or in tropical mountains.............................................................................................. O. sect. Mucidula
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Mycosystema
10
3. Basidiospores subglobose, ellipsoid or amygdaliform, generally smooth, without finger-like ornamentation; widely distributed,
temperate to tropical...................................................................................................................................................... O. sect. Radicatae
3. Basidiospores subglobose, with prominent finger-like ornamentation; Central and South America................O. sect. Dactylosporina
4 DISCUSSION
4.1 Significance of development and hyphal system
of basidioma in relation to systematics of the genus
The manner of basidioma development has widely
been used as a trait in the delimitation of Oudemansiella
and Xerula (Dörfelt 1981; Boekhout & Bas 1986; Pegler
& Young 1987). However, various authors, or even the
same author at different times, differ in their
interpretation of basidoma development in Oudemansiella. For example, Dörfelt (1981, 1982) stated that the
development of O. mucida and O. canarii is hemiangiocarpic (bivelangiocarpic), while the development of
O. radicata is gymnocarpic. Corner (1934) regarded
Collybia apalosarca (= O. canarii) as hemiangiocarpic,
but sixty years later he (Corner 1994) redefined
Oudemansiella as being gymnocarpic with or without a
marginal veil. Our molecular analyses (data not showed)
indicate that sections Mucidula and Oudemansiella have
different ancestors. Thus, even though species in both
sections share similar development patterns, our data
suggest that development in the tropical, subtropical and
south temperate section Oudemansiella and the north
temperate sect. Mucidula is convergent. One explanation
for the strikingly convergent morphologies in the two
sections is that these are structural adaptations to
exposed rotten wood, rather than buried wood as in
sections Radicatae and Dactylosporina. Consequently,
“hemiangiocarpic (bivelangiocarpic) development” or
the formation of a “marginal veil”, i.e., an annulus, may
be an adaptive feature to the environment (Pegler &
Young 1987; Rexer & Kost 1989a).
Some authors used the presence of monomitic or
sarcodimitic structures in the stipititrama (and partially
also in lamellar trama) of Oudemansiella and Xerula as a
criterion for the delimitation of subgenera, genera, or
even family (Boekhout & Bas 1986; Redhead 1987;
Contu 2000). However, whether a stipititrama of
Oudemansiella and Xerula is monomitic or sarcodimitic
appears largely dependant on the judgment and/or
interpretation of the author. For example, Rexer & Kost
(1989a) stated that both O. mucida and O. radicata are
sarcodimitic, while Boekhout & Bas (1986) noted O.
mucida is monomitic and O. radicata is sarcodimitic.
Redhead et al. (1987) stated that both O. mucida and O.
canarii form a tissue somewhat intermediate between
monomitic and sarcodimitic, but that all seven North
American species of Xerula sensu Döferlt (O. sect.
Radicatae in the present sense) have sarcodimitic
stipititramas. In the view point of Corner (1994, 1996),
Oudemansiella is monomitic. As a matter of fact, there
are frequently branched narrow hyphae (“connective
hyphae”) among longitudinally arranged, long-celled,
inflated hyphae (“sarcoskeletals” termed by Boekhout &
Bas, 1986) in the stipititramas of both genera
Oudemansiella and Xerula. However, such “connective
hyphae” are largely restricted to the bulbillate or
ventricose part of the pseudorhiza of species having a
radicating stipe, a similar phenomenon as observed by
Redhead (1987). For the species without a radicating
stipe base, viz. O. mucida, O. canarii, O. orientalis
(Yang 2000), such “connective hyphae” are largely
restricted to the bulbillate or discoid basal part of the
stipe, a portion with only a limited elongation during the
development of the basidioma and the interwoven
arrangement of filamentous or somewhat inflated hyphae
in the disc (Corner 1934) or in the secondary nodulus
(Clémençon 2004) can be traced to certain extent even
when basidioma is mature. In this study, hyphal system
of the stipititrama was not used as a criterion for the
delimitation of genera Oudemansiella and Xerula, or
sections of Oudemansiella.
4.2 Variation of the pileipellis in sections
Oudemansiella and Mucidula
Horak (1968) and Boekhout & Bas (1986)
interpreted the pileipellis of O. platensis and O. canarii
as a cutis. However, Baroni & Ortiz (2002) stated that
the structure of the pileipellis of O. canarii needs further
clarification and illustration. Petersen et al. (2008)
described and illustrated the structures of the pileipellis
Vol.28 No.1
11
and warts or flecks on the pileus of these two species,
sect. Albotomentosae and Xerula sect. Hyalosetae, such
besides O. mucida, in detail.
as O. caussei and O. hongoi (Dörfelt) Zhu L. Yang, and
Our examinations document that the structure of the
X. americana, are not included in Oudemansiella s. str.
pileipellis of O. canarii-O. platensis complex can
as they may form a distinct genus according to our
significantly vary at different stages of development as
preliminary molecular phylogenetic analysis. The unique
suggested by Dörfelt (1981). In young basidioma of O.
ultra-structure of the spore wall of O. nigra and O.
canarii and O. platensis, the pileipellis is an even
xeruloides, the epitunica forming an interrupted and
ixotrichoderm of vertically arranged elements. But with
often discontinuous layer (Pegler & Young 1987),
the expansion of the pileus, the structure of pileipellis in
provides additional sound evidence for the exclusion of
the pileal centre often differs from the pileal margin. In a
these taxa from Oudemansiella.
mature basidioma of O. canarii-O. platensis, the
In our opinion, the eleven taxa of Xerula, with very
pileipellis in the pileal centre remains a ixotrichoderm
detailed and accurate descriptions and illustrations,
with more or less vertically arranged elements (see Figs.
reported from Australia and New Zealand by Petersen
1, 3, and 4) while near the pileal margin it is composed
(2008c) should belong to O. sect. Radicatae, a section
of nearly repent elements. In fully mature or weathered
with diverse taxa widely distributed in many parts of the
basidioma, the pileipellis is often stretched and it
world, rather than sect. Albotomentosae as delimited by
becomes extensively distorted, disrupted, or even
Petersen (2008c).
collapsed, in part due to gelatinization. At this stage it
4.4 Taxa of placement uncertainty
Oudemansiella mediterranea (Pacioni & Lalli) E.
Horak (1988) was initially placed in the genus Hydropus
by Pacioni & Lalli (1985) and then in Flammulina by
Bas & Robich (1988). Quadraccia & Lunghini (1990)
transferred this species to Xerula as X. mediterranea, on
which Xerula subgen. Ixoflammula was based (Contu
2000).
Mycenella kuehneri Romagn., having ornamented
basidiospores, was once placed in Oudemansiella by
Singer (1962b) and, then transferred to the genus Xerula
by Boekhout (1985), Boekhout & Bas (1986) and
Boekhout (1999), but it was retained in the genus
Mycenella by Dörfelt (1985), and Pegler & Young (1987)
based on its overall soft-putrescent habit with a dry
pileus, the non-lignicolous substratum and the absence of
any pseudorhiza. However, Redhead et al. (1987) stated
that the correlated presence of a monomitic stipititrama
and nodose, cyanophylic basidiospores in M. kuehneri
suggests generic distinction.
Molecular phylogenetic data are not available for
either of these taxa. However, the European distribution
of M. kuehneri may suggest that it has no close
relationship with O. steffenii-complex in Central and
South America. Horak (1988) supposed that O.
mediterranea may have affinity with O. caussei and O.
xeruloides, and later treated it as a taxonomic synonym
of O. xeruloides (Horak 2005). Because O. caussei and
looks like a cutis (Dörfelt 1981). A similar phenomenon
was observed in O. venosolamellata (Fig. 6) of sect.
Mucidula. The flecks or warts on the pileus are persistent
(in O. platensis, Fig. 2) or can easily be washed off by
the rain (in O. canarii).
4.3 Excluded groups
Agaricus platyphyllus Pers.: Fr. was placed in the
genus Oudemansiella by Moser (1955) and others.
Knecht (1967), Dörfelt (1981), Lindsey (1986) and
Rexer & Kost (1989a) showed that there are fundamental
differences between Agaricus platyphyllus and others
species of Oudemansiella or Xerula, and should be
regarded as a separate genus, Megacollybia, as proposed
by Kotlaba & Pouzar (1972) and accepted by Pegler &
Young (1987), Redhead (1987), Petersen & Gordon
(1994), and many others.
Pegler & Young (1987) regarded section
Albotomentosae Clémençon as distinct in their system,
which includes O. caussei (Maire) M.M. Moser apud
Clémençon, O. nigra Dörfelt, O. renati Clémençon and
O. xeruloides Bon with O. nigra as the type. The former
three are conspecific according to Boekhout & Bas
(1986), Boekhout (1999) and Contu (2000). In the same
work, they (Pegler & Young 1987) treated Xerula sect.
Hyalosetae Dörfelt, the type of which is X. americana
Dörfelt, as a synonym of O. sect. Xerula Clémençon
typified by Agaricus longipes. Species of Oudemansiella
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Mycosystema
12
O. xeruloides have been excluded from the genus
Oudemansiella in the present paper, the placement of H.
Singapore, 46: 49-75
Corner EJH, 1996. The agaric genera Marasmius, Chaetocalathus,
mediterranea Pacioni & Lalli in Oudemansiella is
Crinipellis, Heimiomyces, Resupinatus, Xerula and Xerulina in
questionable.
Malesia. Beiheft zur Nova Hedwigia, 111: 1-175
Dörfelt H, 1979. Taxonomische Studien in der Gattung Xerula R. Mre.
Acknowledgements: We thank the curators of DUKE, F, FH,
HMAS, HMIGD, HN, MB, MURU, NY, PDD, TENN, TF and
TMI for sending us specimens of Oudemansiella or Xerula on
loan or exchange. Drs. T. Kasuya (Tsukuba), R. Kirschner
(Frankfurt), R.E. Halling (Bronx), Y. Mizuta (Kochi), H. Neda
(Tsukuba), R.H. Petersen (Tennessee), M. Piepenbring
(Frankfurt), S. Garnica (Tuebingen) and S. Takehashi
(Hokkaido) are acknowledged for their various kind help and
Feddes Repertorium, 90: 363-388
Dörfelt H, 1980a. Taxonomische Studien in der Gattung Xerula R. Mre.
(II). Feddes Repertorium, 91: 209-223
Dörfelt H, 1980b. Taxonomische Studien in der Gattung Xerula R. Mre.
(III). Feddes Repertorium, 91: 415-438
Dörfelt H, 1981. Taxonomische Studien in der Gattung Xerula R. Mre.
(V). Feddes Repertorium, 92: 631-674
Dörfelt H, 1982. Die Fruchtkörperentwicklung von Xerula radicata.
Flora, 172: 533-561
Dörfelt H, 1983. Taxonomische Studien in der Gattung Xerula R. Mre.
support to this study.
(VII). Feddes Repertorium, 94: 251-62
Dörfelt H, 1984. Taxonomische Studien in der Gattung Xerula R. Mre.
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